A tentative classification of the palearctic unionids

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ZOOLOGICAL SERIES

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UNIVERSITY OF ILLINOIS
FIELD MUSEUM OF NATURAL HISTORY

Volume 24 CHICAGO, JANUARY 30, 1940 No. 11

A TENTATIVE CLASSIFICATION OF THE
PALEARCTIC UNIONIDS

BY FRITZ HAAS

CURATOR OP LOWER INVERTEBRATES

INTRODUCTION

The arrangement which is to follow is based upon more than
thirty years' experience in work on unionids in general and on the
palearctic ones in particular; it aims to place the pearly fresh-water
mussels of the palearctic region within the general system of the
unionids, and to show the natural inter-relationships of the forms.
This requires some mention of earlier taxonomic essays on the subject.

Until the middle of the last century, that is to say, until the time
of Rossmaessler in Germany and Dupuy in France, Linnaean
methods prevailed; every animal believed to be unknown to science
was described as a new species and nobody worried about the
natural relations of the "species" thus originated. This being so, it
is not surprising to find that many phaenotypic features were mis-
taken for specific ones and that many forms which had nothing to
do with each other were thrown together. It seems that Rossmaessler
was the first to recognize clearly the transformation of the unionid
shell by environmental conditions; he, at least, knew about the
shaping influence of lacustrine life, which manifests itself in the
larvation of the still specifically characteristic juvenile shell, or in
that of the fluviatile phase of unionids.

The following era may be characterized by the methods of J. R.
Bourguignat and his pupils — Locard, Servain, Coutagne, and many
others. This nouvelle ecole attributed specific value to all the differ-
ences of shape in fresh-water mussels, which may have originated
through environmental influences as well as through geographic
isolation. Under this school, discrimination went far more into
detail than under the Linnaean method: each shell which differed
from an already known mussel in three dimensions, or by the indices
calculated from them, was a new species. It is obvious that such a

No. 464 115

116 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

method, because of the impossibility of securing exact measurements
in the always somewhat ventricose unionid shell, could afford
approximate figures only; it led inevitably to the description of
every individual examined as a new species. Umbonal sculpture
and details of the hinge composition were not considered at all.
Thus, convergencies due merely to similarity of measurements in
the three dimensions, often led to unwarranted inferences with
regard to natural relations. The placing of Margariti/era margariti-
fera and of Psilunio littoralis close to certain elongated forms of
crassoid unios affords an example.

Truth compels us to admit that, leaving aside slips of judgment
due to deficiencies of the method, the natural arrangement of the
many described "species" was not spoiled; on the contrary, they
generally were united into groups, which correspond to a certain
degree to our modern racial groups. The geographical factor, how-
ever, without which a natural taxonomic arrangement can not now
be imagined, does not play any role in the Bourguignatian method,
nor was the anatomy of the soft parts considered.

Another Frenchman, H. Droue't, stands quite alone among his
contemporaries, by reason of the emphasis he placed upon the con-
nection between the shape of his new unionids and the life conditions
of their respective habitats. Droue't was, indeed, a pioneer, but he
knew too little about the ecology of fresh-water mussels and was
therefore often deceived by convergent shapes.

After Rossmaessler's death, at a time when descriptions of new
unionids were produced by the hundreds in France, in Italy, and in
other countries, work on fresh-water mussels lagged in Germany.
The German malacologists (especially Kobelt, who continued Ross-
maessler's incomplete Iconographie} did not agree with Bourguignat's
views, which were also in disfavor with some of the French spe-
cialists; for example, with the editors and collaborators of the
Journal de Conchy 'liologie. Kobelt's disapproving attitude had won
him Bourguignat's intense hatred, but he kept to his own ideas on
unionids, which he did not publish until after Bourguignat's death.

The list of the Unionidae in C. A. Westerlund's Fauna der in der
palaarctischen Region lebenden Binnenconchylien (7, 1890) shows
how Bourguignat and his school, to which Westerlund belonged,
believed the fauna of palearctic fresh-water mussels to be composed.

More modern ideas on this subject originated with Kobelt, who,
in various publications, pointed out the isolating effect of hydro-
graphic frontiers. By many examples he tried to prove the theory,

THE LIBRARY OF THE

1940 PALEARCTIC UNIONIDS — HAAS FFR1^?) 1QAH

upon which his principal conclusion was based, that a^llRSffi&Pf ILLINOIS

arrangement of the unionids had to take into consideration their

distribution in the different hydrographic systems. When this view

was accepted, and it was understood that the overwhelming num-

jber of described palearctic unionids must be reduced to a very

ol limited number of "fundamental" species, there was the unfortunate

0 circumstance that nobody knew which these fundamental species
gmight be.

Everyone agreed, at least, that these fundamental species, which-
Oever they might be, had given origin to local races in the different
^ parts of the river systems inhabited by them. Collectors and scien-
cotists — among them, thirty years ago, even the writer of these
"-'lines — began to prove the existence of these local races and to
<describe them wherever necessary. In Germany, where the Bour-
sguignat method had not been so popular as it had been in France,
Italy, and the lower Danube Basin, comparatively few unionids
had been described, aside from the fundamental ones, so that there
were few names available for such new races as seemed to be worth
describing. New names were invented, and I plead guilty to having
encumbered nomenclature with not less than fifteen new denomina-
tions, bestowed mostly on mussels from western Germany.

Looking backward, I can not wholly disapprove of my course
at that time. I have come to realize that the differences mistaken
by me for racial features were phaenotypic, due to environmental
factors; furthermore, I have learned that the limited geographic
areas inhabited by such uniformly characterized fresh-water mussels,
<o were nothing but areas of uniform conditions of life. But even if

- such distinct local forms, for instance, those of the Unio crassus
group, are not worth retaining as trinomially named races, they
are nevertheless somewhat more than mere variations. They can
be considered to be incipient races, having a mutative basis. That

1 such slight differences, which are nevertheless distinguishable to an
experienced eye, can not be made the basis of named forms, is due

- to our present system of nomenclature, which does not go beyond
. a third name. I have been greatly pleased to learn from modern

herpetologists and ornithologists that they have found similar con-
ditions and feel some need for a method of designation for the
above-mentioned "subraces," which we may call incipient ones. I
can not but agree that this would be desirable.1

1 William Morton Wheeler, in his work on ants, consistently employs a quad-
rinomial nomenclature; cf., for example, his Ants (1910).

118 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

We shall leave aside, as being far from our present object, the
merely geographical part of the unionid problem, which, following
Kobelt's ideas, involved the tracing of old and bygone connections
between now separated river systems, and other related subjects.

The arrangement of the palearctic unionids which follows rests
mostly on Kobelt's ideology. It is backed by long experience in
field observation and indoor study and by personal knowledge of
many types of unionids contained in the extremely rich collection
of the Senckenberg Museum in Frankfort-on-the-Main or entrusted
to me for study by other museums; for example, that of Geneva,
Switzerland, which possesses Bourguignat's types, and the United
States National Museum, where I have had the privilege of
studying Lea's types of unionids from Asia Minor. The investiga-
tion of such authentic material often gave results quite different
from those derived from the study of descriptions and figures; the
attributions to synonymy to be found in my arrangement, therefore,
often differ from those customary, and are the immediate conse-
quence of personal acquaintance with the respective "species."

As to the position of the palearctic unionids within the general
system of the family, my arrangement also differs from that of
other writers, and even from that of Thiele in his Handbuch der syste-
matischen Weichtierkunde. In many cases I have no anatomical
evidence, and I trust that my system may be attributed to special
acquaintance with the subject; paleontological data have frequently
influenced my ideas upon the relationships between palearctic and
exotic genera.

The following arrangement is only a tentative one. In France,
where Bourguignat's influence prevailed up to the beginning of this
century, Germain in his Mollusques terrestres et fluviatiles de la France
(22, Part 2: Faune de France, Paris, 1931) reduces the number of
French unionids to a very few fundamental species. It is thus
evident that the endeavor to simplify and to concentrate the system
is everywhere obvious. Since others have ideas similar to mine,
they may perhaps welcome my arrangement as a basis for future
and more extensive research.

Finally, I must try to explain my concept of a "subspecies" or
a "local race," in order to justify my arrangement. I realize that,
in many cases, it is very difficult to distinguish two neighboring
local races, as they seemingly are linked by intermediate forms.
The impossibility of a sharp separation may be due to overlappings
at the borders of distribution or, possibly, to hybridizations; but in all

1940 PALEARCTIC UNIONIDS— HAAS 119

such cases, the respective subspecies are more easily distinguishable
at the centers than at the borders of their areas. Something of the
same kind, only on a much smaller scale, also happens to adjacent full
species; for instance, to Unio pictorum Linnaeus and Unio elongatulus
C. Pfeiffer, which can scarcely be distinguished in eastern France,
where there are certainly hybridizations between them.

In cases such as these, the distinction between species or sub-
species is very difficult, and I have usually based my decisions upon
the shape of the shells found in the centers of the areas of distribution
in question. I am well aware of the tentative character of such an
arrangement; larger series of material from new localities may
necessitate much revision of the system set forth in this paper.

The present paper recognizes 9 genera, 19 species, and 61 sub-
species of palearctic unionids.

TENTATIVE ARRANGEMENT OF THE PALEARCTIC
UNIONIDS INTO NATURAL GROUPS

Family Margaritiferidae

The palearctic margaritiferids all belong to the genus Margari-
tifera Schumacher; the following species or specific groups are known:

Genus Margaritifera Schumacher, 1816

Margaritana Schumacher, 1817 Pseudunio Haas, 1910

Potamida Swainson, 1840

Margaritana syriaca Pallary, 1929, is clearly a Psilunio. See
under Psilunio littoralis homsensis Lea, page 135.

Margaritifera auricularia Spengler, 1793
Margaritifera auricularia auricularia Spengler, 1793

Unio sinuatus Lamarck, 1819 Unio margaritanopsis Locard, 1889

Iberian Peninsula; France; Po basin in North Italy. Subfossil in
England (Thames) ; Germany (Thuringia, Rhine basin) ; central Italy.

Margaritifera auricularia maroccana Pallary, 1920

Margaritana redomica Pallary, 1927 Margaritana dernaica Pallary, 1928

Very closely related to typical auricularia, but probably dis-
tinguishable as a local race.
Morocco.

120 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Margaritifera margaritifera Linnaeus, 1758

A holarctic species, represented in the palearctic region by the
following four subspecies:

Margaritifera margaritifera margaritifera Linnaeus, 1758

Unio elongata Lamarck, 1819 Margaritana pyrenaica Bourgui-

Unio roissyi Michaud, 1831 gnat, 1889

Unio margaritifer var. minor Ross- Margaritana alleni Castro in Lo-

maessler, 1835 card, 1889

Unio brunneus Bonhomme, 1840 Margaritana margaritifera parvula

Unio tristis Morelet, 1845 Haas, 1908

Margaritana freytagi Kobelt, 1886 Margaritana durrowensis Phillips,

Margaritana michaudi Locard, 1889 1928

Part of the names quoted in the synonymy designate incipient
races which have no status in modern nomenclature.

Extra-mediterranean Europe.

Margaritifera margaritifera dahurica Middendorff, 1851
Eastern Siberia.

Margaritifera margaritifera middendorffi Rosen, 1926

Unio complanatus Middendorff (not Solander) 1851

Kamchatka.
Margaritifera margaritifera laevis Haas, 1910

Margaritana dahurica Kobelt (not Margaritana sachalinensis Shadin,

Middendorff), 1879 1938

Ptychorhynchus laevis Haas, 1910

Saghalin; northern Japan.

Family Unionidae
Subfamily Unioninae

Some of the genera grouped here, such as Leguminaia, Pseudo-
dontopsis and Microcondylaea, the anatomy of which is entirely
unknown or only partly known, may not belong to this subfamily
at all.

Genus Unio Retzius, 1788

Palearctic, entering into the Oriental region just in the East
and the Southeast; closely related to the nearctic genus Elliptic
Rafinesque.

Unio pictorum Group

Very similar and probably related to the East-Asiatic Unio
douglasiae Griffith and Pidgeon and its group.

1940

PALEARCTIC UNIONIDS — HAAS

121

Unio pictorum Linnaeus, 1758

Palearctic region, except the Mediterranean basin, where it is
found only in the Rhone system, in North Africa and in the rivers
flowing into the Black Sea. In the parts of the Mediterranean basin
where it is absent, it is represented by the races of Unio elongatulus
C. Pfeiffer. In the Rhone system as well as in North Africa, both
species occur side by side; in central France (Rhone basin and rivers
of the central plain) they apparently hybridize, for the unios of the
pictorum group are intermediate between these two forms and are
distinguished by extreme variability.

Unio pictorum pictorum Linnaeus, 1757

Unio limosus Nilsson, 1882
Unio deshayesi Michaud, 1832
Unio pictorum var. grandis Ross-

maessler, 1842
Unio maltzani Ktister, 1854
Unio pictorum var. niger Joannis,

1858

Unio platyodon Jordan, 1879
Unio jousseaumi Bourguignat, 1882
Unio pincianus Bourguignat, 1882
Unio dolfussianus Bourguignat,

1882

Unio joannisi Bourguignat, 1882
Unio gallicus Bourguignat, 1882
Unio cancrorum Bourguignat, 1882
Unio rhynchetinus Letourneux, 1882
Unio rostratellus Bourguignat, 1882

Unio torsatellus Berthier, 1882
Unio hollandrei Bourguignat, 1882
Unio fascellinus Servain, 1882
Unio f alms Bourguignat, 1882
Unio strigatus Servain, 1886
Unio caravellus Servain, 1887
Unio lesumicus Bourguignat, 1888
Unio subbalatonicus Servain, 1888
Unio brebissoni Locard, 1889
Unio campylus Locard, 1889
Unio siliquiformis Locard, 1889
Unio macropisthus Bourguignat,

1889

Unio hospitali Locard, 1889
Unio oesiacus Locard, 1889
Unio rectus Locard, 1890
Unio battonensis Kobelt, 1896

The synonymy of this and the following races is still very much
confused, especially for the French forms. It was due to their vari-
ability that so many "species" and "varieties" were described in
France. They could not be interpreted by Germain (1931). Hence
the list of synonyms is neither correct nor complete.

Atlantic northern and central and northwestern Europe; appar-
ently hybridizing with Unio elongatulus C. Pfeiffer in eastern France.

Unio pictorum praeposterus Kiister, 1854
Naab; Regen; Thaya (Danube system).

Unio pictorum latirostris Kiister, 1854

Unio concinnus Kiister, 1840 (no- Unio baletonicus Ktister, 1861

men nudum) Unio balatonicus Servain, 1881

Unio quinqueannulatus Krister, 1854 Unio dubreuili Servain, 1881

Unio area Held in Ktister, 1854 Unio eucallistus Kobelt, 1915
Unio decollatus Held in Ktister, 1854

Haas and Schwarz (1913) regard U. quinqueannulatus as a hybrid
between U. pictorum latirostris and its northern neighbor U. pictorum
pictorum.

German Danube; Hungary; entering into the Save system.

122 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24
Unio pictorum platyrhynchus Rossmaessler, 1835

Unio longirostris Rossmaessler, 1836 Unio graniger J. F. Schmidt, 1847

Unio ponderosus Spitzi in Ross-
maessler, 1842

Styria; Carinthia; Carniolia (Danube system).

Unio pictorum middendorffii Westerlund, 1890
Eastern Siberia, still in the Amur River.

Unio pictorum schrenckianus Clessin, 1880

Unio pictorum var. okae Kobelt, Unio pictorum var. okensis Shadin,

1911 1938

Unio annulatus Kobelt, 1912 Unippictorum var. pygmaeus Sha-

Unio pictorum schrenckianus var. din, 1938

dicki Model!, 1930 Unio pictorum var. defectivus Sha-

Unio lindholmi Shadin, 1938 din, 1938

Pontic Russia.

j

Unio pictorum ascanius Kobelt, 1913
Northern Asia Minor.

Unio pictorum proechistus Bourguignat, 1870
Rumanian Danube; Vardar River (?)

Unio pictorum gaudioni Drouet, 1881

Unio gentilis Haas, 1911

Doiran Lake, Macedonia; Maritza River; Varna, Bulgaria;
vicinity of Istanbul.

(?)Unio pictorum proechus Bourguignat, 1862

Unio actephilus Bourguignat, 1862

Lake Lucerne, Switzerland.

(?)Unio pictorum moussonianus Clessin, 1887
Lake Zug, Switzerland.

Unio pictorum rostratus Lamarck, 1819

The synonymy still being unsettled, the compilation is tentative:

Unio pictorum var. vinceleus Joan- Unio lugdunicus Coutagne, 1889

nis, 1858 Unio cristulatus Drouet, 1889

Unio pictorum var. lumens Joannis, Unio perroudi Locard, 1889

1858 Unio hydrellus Locard, 1889

Unio gestroianus Bourguignat, 1882 Unio bramicus Baichere, 1890

Unio charpyi Drouet, 1888

Occasionally hybridizing with Unio elongatulus C. Pfeiffer, where
they occur together.

Rhone system, France.

1940

PALEARCTIC UNIONIDS — HAAS

123

Unio pictorum platyrhynchoideus Dupuy, 1849

(l)Unio michaudiana Des Moulins,
1833

Unio requienii var. crassidens Ross-
maessler, 1844 (not Lamarck,
1819)

Unio philippei Dupuy, 1849

Unio danielis Gassies, 1867

Unio moreleti Folin and Berillon,
1874 (not Deshayes, 1848)

Unio brindosianus Folin and Beril-
lon, 1874

Unio baudoni Folin, 1874

Unio moreletianus Folin and Beril-
lon, 1877

Unio moriscottei Folin, 1877
Unio bayonnensis Folin and Beril-
lon, 1877

Unio berilloni Locard, 1882
Unio hauterivianus Bourguignat,

1882

Unio corbini Bourguignat, 1882
Unio milne-edwardsi Bourguignat,

1882
Unio fagoti Bourguignat, 1883

Southwestern Atlantic France, including the Garonne River.

Unio pictorum mucidus Morelet, 1845

Unio nevesi Castro, 1885
Unio simoesi Castro, 1885
Unio camera tus Drouet, 1893
Unio limosellus Droue't, 1893
Unio decurtatus Droue't, 1893
Unio callipygus Drouet, 1893
Unio aeschrus Locard, 1899
Unio tameganus Locard, 1899
Unio onconensis Locard, 1899
Unio hypoxanthus Locard, 1899
Unio chorellus Locard, 1899
Unio chorellinus Locard, 1899
Unio submucidus Locard, 1899
Unio barbosanus Locard, 1899
Unio castroi Locard, 1899
Unio silvai Locard, 1899

Unio ocresanus Locard, 1899
Unio paulinoi Locard, 1899
Unio subhispanus Locard, 1899
Unio hyperephanus Locard, 1899
Unio chasmirhynchus Locard, 1899
Unio mundanus Locard, 1899
Unio euchasmus Locard, 1899
Unio schousboei Locard, 1899
Unio taganus Locard, 1899
Unio abrantesianus Locard, 1899
Unio scalabisianus Locard, 1899
Unio allenianus Locard, 1899
Unio cyrtus Locard, 1899
Unio sousanus Locard, 1899
Unio novus Locard, 1899
Unio neothaumus Locard, 1899

Northwestern part of Atlantic Iberian Peninsula.
Unio pictorum delphinus Spengler, 1793

Unio gibbus Spengler, 1793
Unio hispanus Rossmaessler, 1844
Unio dactylus Morelet, 1845
Unio lusitanus Drouet, 1879
Unio (hispanus var.?) sevillensis
Kobelt, 1887

Unio hispanus var. sphenoides Wes-

terlund, 1892

C!)Unio gravatus Drouet, 1893
Unio turdetanus Drouet, 1893
Unio taginus Kobelt, 1903

Southwestern part of Atlantic Iberian Peninsula.
Unio pictorum ravoisieri Deshayes, 1848

Unio atharsus Bourguignat, 1889

Mediterranean North Africa, including Tunisia in the east.
Further synonymy still unsettled.

Unio tigridis Bourguignat, 1852
Southwestern Asia.

124 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Unio tigridis tigridis Bourguignat, 1852

Unio truncatus Swainson, 1829 (not

Spengler, 1793)
Unio hueti Bourguignat, 1855
Unio natolicus Kiister, 1856
Unio mussolianus Kiister, 1861
Unio rasus Lea, 1863
Unio mosulensis Lea, 1863
Unio bourguignatianus Lea, 1863
Unio dignatus Lea, 1863
Unio delicatus Lea, 1863
Unio tigris Lea, 1870
Unio anemprosthus Locard, 1883
Unio subtigridis Locard, 1883
Unio axiacus Locard, 1883
Unio chantrei Locard, 1883

Syria, in the Orontes basin; Iraq; Lake Urmiah.
Unio tigridis terminalis Bourguignat, 1852

Uniojouberti Locard, 1883
Unio antiochianus Locard, 1883
Unio kobelti Rolle, 1895
Unio tigridis var. harunis Kobelt,

1912
Unio dignatus semiramidis Kobelt,

1913
Unio dignatus assuricus Kobelt,

1913

Unio dignatus ninusi Kobelt, 1913
Unio hyperamblius Kobelt, 1913
Unio calliopsis Kobelt, 1913
Unio siouffi Kobelt, 1913
Unio schwarzii Kobelt, 1915

Unio grelloisianus Bourguignat,

1856
Unio grelloisianus var. giganteus

Bourguignat, 1856
Unio lunulifer Bourguignat, 1856
Unio jordanicus Bourguignat, 1865
Unio pietri Locard, 1880
Unio lorteti Locard, 1880

Unio tristrami Locard, 1883
Unio ellipsoideus Locard, 1883
Unio genezarethanus Locard, 1883
Unio tiberiadensis Locard, 1883
Unio -prosacrus Locard, 1883
Unio zabulonicus Locard, 1883
Unio her odes Kobelt, 1895
Unio kisonis Kobelt and Rolle, 1895

For this subspecies, Prashad (1919) has introduced the sub-
generic name of Eolymnium, apparently without sufficient reason.
Littoral rivers of Cilicia, Syria and Palestine; Jordan River.

Unio abyssinicus von Martens, 1866

]Unio vnllcocksi Newton, 1899
\Unio vignardi Pallary, 1924

\Unio fayumensis Pilsbry and Be-
quaert, 1927

Abyssinia; the subfossil forms (t) in the basin of the Lower Nile.

Unio elongatulus Group

Unio elongatulus C. Pfeiffer, 1825
Mediterranean region; Asia Minor.

Unio elongatulus elongatulus C. Pfeiffer, 1825

Unio lijacensis Kobelt, 1890 Unio lijacensis var. gallensteini

Kobelt, 1890

Area north of the Adriatic Sea: Tagliamento River(?); basin of
Isonzo River; northern Dalmatia(?)

Unio elongatulus pallens Rossmaessler, 1842

Unio viridiflavus Kiister, 1854 Unio petrovichi Kiister, 1854

Unio petterdianus Kiister, 1854 Unio nitidosus Drouet, 1879

Central and southern Dalmatia (Narenta, Imoshi); north Greece;
River Save in Croatia (?)

1940 PALEARCTIC UNIONIDS— HAAS 125

Unio elongatulus fiscallianus Kleciach, 1872

Imoshi, Dalmatia; may turn out to be only a lacustrine form of
U. elongatulus pollens Rossmaessler.

Unio elongatulus sandrii Rossmaessler, 1844

Unio nuperus Ziegler, in litt. Unio dalmaticus Drouet, 1879

Unio sandrii var. sericatus Ross- Unio succineus Drouet, 1881

maessler, 1844 Unio morlachicus Kobelt, 1915

Unio ceratinus Drouet, 1879

Northern Dalmatia; may be identical with Unio elongatulus
pallens Rossmaessler.

Unio elongatulus decipiens Drouet, 1881

Dalmatia; Montenegro. Possibly a lacustrine form of Unio
elongatulus pallens Rossmaessler.

Unio elongatulus kriiperi Drouet, 1879

Unio destructilis Villa in Kobelt, 1915

Montenegro; Lake Vrachori in Aetolia, Greece. This race may
be only a degenerate form of Unio elongatulus pallens Rossmaessler.

Unio elongatulus quelleneci Drouet, 1895

Unio copaisanus Drouet, 1895

Lake Kopais, Aetolia, Greece; possibly a lacustrine form of Unio
elongatulus pallens Rossmaessler.

Unio elongatulus bourgeticus Bourguignat, 1882

Unio sabaudianus Bourguignat, Unio mucidellus Bourguignat, 1889

1882 Unio voltzii Kobelt, 1911

Unio riciacensis Bourguignat, 1882 Unio voltzii var. ursannensis Ko-

Unio orthus Coutagne, 1882 belt, 1911
Unio occidaneus Drouet, 1888

Basin of River Saone in western France; Swiss Jura; Savoy;
introduced into the Rhine system (Miihlhausen) by means of the
Rhine-Rhone Canal.

A very difficult and until recently little understood race, often
confounded with forms of Unio crassus Retzius on account of its
oval shape and wavy umbonal sculpture. The above list of synonyms
is provisional.

Unio elongatulus turtoni Payraudeau, 1826

Unio capigliolo Payri
Unio bandini Klister,

Corsica, Sardinia.

Unio capigliolo Payraudeau, 1826 Unio exauratus Locard, 1889

Unio bandini Klister, 1837

126 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Unio elongatulus moquinianus Dupuy, 1843

Unio antimoquinianus Locard, 1889

Unio consentaneus var. moquini
Germain, 1931

Hautes-Pyre'ne'es; Basses-Pyre'ne'es; southern France.

This race has also often been confounded with forms of crassus.
The synonymy will certainly prove to be more extensive.

Unio elongatulus aleroni Companyo and Massot, 1865

Central southern France (HeYault, Aude, Pyre'ne'es-Orientales,
Tarn, Haute-Garonne, Ariege).

Unio elongatulus rousii Dupuy, 1849

Unio thermalis Dupuy, in litt.

Unio malafossianus Bourguignat,

1882

Southern France (Garonne, Gers). Often confounded with Unio
tumidus Retzius; synonymy still incompletely known.

Unio elongatulus niancus Lamarck, 1819

Unio ardiisianus Reynies, 1843
Unio condatinus Letourneux, 1882
Unio mucidulus Bourguignat, 1882
Unio gobionum Bourguignat, 1882
Unio mongazonae Servain, 1887
Unio asticus Servain, 1887

Unio eutrapelus Servain, 1887
Unio oberthurianus Bourguignat,

1889

Unio arcuatulus Bourguignat, 1889
Unio amblyus Castro, 1889

A very variable, difficult form, often confounded with forms of
crassus; occasionally hybridizes with Unio pictorum Linnaeus.

Central France, mainly basins of rivers Seine and Marne.
Unio elongatulus requienii Michaud, 1831

Unio jacquemini Dupuy, 1849
Unio saintsimonianus Fagot, 1882
Unio veillanensis Blanc, 1882
Unio orthellus Berenguier, 1882
Unio jourdheuili Ray, 1882
Unio forojuliensis Berenguier, 1882
Unio triffoiricus Bourguignat, 1885
Uniofabaeformis Bourguignat, 1889
Unipfrayssianus : Bourguignat, 1889
Unio meyrannicus Bourguignat,
1889

Further synonymy still unsettled, the above list not final.
Southeastern part of France, the Rhone, and east of the Rhone.

Unio elongatulus penchinatianus Bourguignat, 1865

Rivers of Mediterranean coast of northeastern Spain, to and
including the Ebro.

Unio aramonensis Locard, 1889
Unio vardonicus Locard, 1889
Unio talus Bourguignat, 1889
Unio souzanus Castro, 1889
Unio royanus Locard, 1889
Unio aegericus Locard, 1889
Unio mucidulinus Locard, 1889
Unio baicheri Locard, 1890
Unio arelatus Bourguignat, 1892

1940

PALEARCTIC UNIONIDS — HAAS

127

Unio elongatulus valentinus Rossmaessler, 1854

Unio graellsianus Bourguignat, Unio courquinianus Bourguignat,

1865 1865

Unio almenarensis Drouet, 1893

Rivers of Mediterranean coast of Spain, south of the Ebro;
southern border unknown.

Unio elongatulus gargottae Philippi, 1836

(l)Unio lobata Jan, 1832
Unio aradae Philippi, 1844
Unio aradasii Kobelt, 1876
Unio benoiti Bourguignat, 1883
Unio monterosati Bourguignat, 1883
Unio bivonianus Bourguignat, 1883
Unio caficianus Bourguignat, 1883

Sicily.

Unio biformis Monterosato, 1896
Unio bitortus Monterosato, 1896
Unio bipartitus Monterosato, 1896
Unio ricconianus Monterosato, 1896
Unio cuspidatus Monterosato, 1896
Unio subindentatus Adami in Mon-
terosato, 1896

Unio elongatulus lawleyanus Gentiluomo, 1868

Unio larderelianus Pecchioli, 1869
Unio romanus Kobelt, 1875
Unio cumensis Kobelt, 1880
Unio cumanus Kobelt, 1880
Unio pornae Bourguignat in Lo-

card, 1880
Unio meretricis Bourguignat in

Locard, 1880

Unio moltenii Adami, 1882
Unio umbricus Adami, 1882
Unio etruscus Drouet, 1883
Unio campanus Drouet, 1883

Peninsular Italy.

Unio meridionalis Drouet, 1883
Unio latinus Bourguignat, 1883
Unio campsus Bourguignat, 1883
Unio uziellii Bourguignat, 1883
Unio pisanus Bourguignat, 1883
Unio gentiluomoi Bourguignat,

1883

Unio pecchiolii Bourguignat, 1883
Unio polii Drouet, 1883
Unio planci Drouet, 1883
Unio isseli Bourguignat, 1883
Unio d'Anconae Bourguignat, 1883

Unio elongatulus glaucinus Porro, 1838

Unio corrosus Villa, 1841
Unio spinellii Villa, 1852
Unio requienii var. vulgaris Stabile,

1859
Unio requienii var. oriliensis Stabile,

1859
Unio requienii var. blauneri Stabile,

1859

Unio gestroianus Bourguignat, 1871
Unio villae Issel, 1871
Unio larius Drouet, 1879
Unio gurkensis Bourguignat, 1881
Unio brachyrhynchus Drouet, 1881
Unio opisodartos Adami, 1882
Unio stephaninii Adami, 1882
Unio subcylindricus Drouet, 1883
Unio fluminalis Drouet, 1883
Unio delpretei Bourguignat, 1883
Unio gredleri Drouet, 1883
Unio minusculus Drouet, 1883
Unio benacinus Drouet, 1883
Unio siliquatus Drouet, 1883
Unio nitidus Drouet, 1883

Unio eucallistellus Bourguignat,

1883

Unio callichrous Bourguignat, 1883
Unio padanus Bourguignat, 1883
Unio strobeli Bourguignat, 1883
Unio pedemontanus Bourguignat,

1883

Unio longobardus Drouet, 1883
Unio brianteus Drouet, 1883
Unio sabiniensis Bourguignat, 1883
Unio verbanicus Bourguignat, 1883
Unio athesinus Adams, 1885
Unio ruffonii Kobelt, 1886
Unio humerosus Westerlund, 1890
Unio humerosus var. costanus Tie-

senhausen, 1894
Unio directm Drouet, 1895
Unio palustris Drouet, 1895
Unio humerosus var. elongata Ko-
belt, 1915
Unio humerosus var. roboretana

Kobelt, 1915
Unio destructilis Villa in Kobelt, 1915

Basin of the Po, and Adige, continental Italy.

128 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24
Unio elongatulus eucirrus Bourguignat, 1857

Unio raymondi Locard, 1883
Unio alexandri Kobelt and Rolle,

1895
Unio cilicicus Kobelt and Rolle,

1895
Unio cilicicus var. adanensis Rolle,

1895
Unio cilicicus var. • jenemterensis

Kobelt and Rolle, 1895

Unio cilicicus var. subsaccatus Ko-
belt and Rolle, 1895
Unio berythensis Kobelt and Rolle,

1895

Unio raymondopsis Kobelt, 1913
Unio kuweikensis Kobelt, 1913
Unio sesirmensis Kobelt, 1913
Unio (cilicicus var.) anceyi Kobelt,
1913

Littoral rivers of Cilicia, Syria, and Palestine.
Unio elongatulus dembeae Sowerby, 1865

Unio aeneus Jickeli, 1874
Unio jickelii Bourguignat, 1883
Unio alfierianus Bourguignat, 1885
Unio hamyanus Bourguignat, 1885
Unio ilqi Bourguignat, 1885

Abyssinia.

Unio meneliki Bourguignat, 1885
Unio soleilleti Bourguignat, 1885
Unio traversii Pollonera, 1888
Unio erlangeri Kobelt, 1909

Unio elongatulus durieui Deshayes, 1847

(I) Unio emarginatus Lea, 1834
Unio tafnanus Kobelt, 1884
Unio sitifensis Kobelt, 1884
Unio issericus Kobelt, 1884
Unio tetuanensis Kobelt, 1884

Unio medjerdae Kobelt, 1886
Unio micelii Kobelt, 1886
Unio delevieleusae Hagenmiiller,
1887

After studying the type of Unio emarginatus Lea (U. S. Nat.
Mus. No. 86159) I can not decide whether this form, known only by
this type, is identical with durieui or not; the type specimen is com-
posed of two odd valves.

Mediterranean North Africa.

Unio tumidus Group
Unio tumidus Retzius, 1788

Atlantic and Pontic parts of Europe.

Unio tumidus tumidus Retzius, 1788

Unio conus Spengler, 1793

Mya depressa Donovan, 1802

Mya ovata Donovan, 1802

Mya ovalis Montagu, 1803

Ml/sea solida Turton, 1822

Unio inflata Hecart, 1833

Unio muUeri Rossmaessler, 1838

Unio tumidus var. saccatus Ross-
maessler, 1854

Unio pictorum var. niger Joannis,
1859

Unio tumidus var. pictus Beck in
Moerch, 1864

Unio tumidus var. maxima Moerch,
1864

Unio tumidus var. limicola Moerch,
1864

Unio batatms var. incurvatus Col-
beau, 1868

Unio tumidus var. heckingi Colbeau,
1868

Unio tumidus var. rohrmanni Ko-
belt, 1880

Unio bardus Bourguignat, 1881

Unio edyus Bourguignat, 1882

Unio alpecanus Bourguignat, 1882

1940

PALEARCTIC UNIONIDS — HAAS

129

Unio fourneli Bourguignat, 1882
Unio mulierum Servain, 1882
Unio schroederi Bourguignat, 1885
Unio rhenanus Kobelt, 1886
Unio tumidus var. godetiana Cles-
sin, 1887
Unio tumidulus Locard, 1889

Unio aldemaricus Bourguignat, 1889
Unio giberti Locard, 1889
Unio rothomagensis Locard, 1890
Unio albovicensis Locard, 1893
Unio lacrymiformis Locard, 1893
Unio levoiturieri Locard, 1893
Unio lauterborni Haas, 1909

Even the brief synonymy of this widespread form is still un-
settled, since it is known that it has often been confounded with
short, stout forms of Unio pictorum Linnaeus.

Atlantic western and central Europe.
Unio tumidus borysthenensis Kobelt, 1880

Unio gerstfeldtianus Clessin, 1880
Unio tumidus var.falcatulus Drouet,

1881

Unio tumidus ilekensis Kobelt, 1912
Unio tumidus var. natermanni Ho-

nigmann, 1914
Unio tumidus var. bashkiricus

Shadin, 1938

Unio tumidus var. moltshanovi

Shadin, 1938
Unio tumidus var. fridmani Shadin,

1938
Unio kobeltianus Shadin, 1938 (not

Plaas, 1913)

European streams draining into the Black Sea: lower part of
Danube system; southern Russia.

Unio crassus Group

The members of this group have often been confounded with
elongated forms of Psilunio and with round forms of Unio elongatulus.

Unio crassus Retzius, 1788

Unio crassus crassus Retzius, 1788

Unio musivus Spengler, 1793

Unio ater Nilsson, 1822

Unio rubens Menke, 1828

Unio rugatus Menke, 1828

Unio crassus var. maximus Kobelt,

1872

Unio pseudolittoralis Clessin, 1875
Unio kochi Kobelt, 1886
Unio hamburgiensis Servain, 1888

Unio pseudolittoralis var. curonicus

Riemenschneider, 1907
Unio crassus heimburgi Haas, 1911
Unio crassus thuringiacus Israel,

1917
Unio crassus polonicus Polinski,

1917

Unio crassus ornatus Polinski, 1917
Unio crassus var. ishnensis Shadin,

1938

North Europe; Atlantic Central Europe, in the west to the
Rhine, but not in it; Atlantic and boreal Russia.

Unio crassus cytherea Kiister, 1833

(l)Unw planus Studer, 1820; un-
identifiable

Unio sinuatus Studer, 1820 (nomen
nudum)

Unio dilatatus Studer, 1820 (nomen
nudum)

Unio ovatus Studer, 1820 (nomen

nudum)

Unio sinuolata Kiister, 1833
Unio dubius Fitzinger, 1833 (nomen

nudum)

130 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Unio nigricans Fitzinger, 1833

(nomen nudum)
Unio castaneus Fitzinger, 1833

(nomen nudum)

Unio decurvatus Rossmaessler, 1835
Unio labacensis Rossmaessler, 1835
Unio vitreus Rossmaessler, 1835
Unio atrovirens Rossmaessler, 1836
Unio amnicus Rossmaessler, 1836
Unio piscinalis Rossmaessler, 1836
Unio reniformis Rossmaessler, 1836
Unio consentaneus Rossmaessler,

1836

(1}Unio retusa Held, 1836
Unio batavus var. gibbosus Held,

1836
Unio batavus var. elongatus Held,

1836
Unio carinthiacus Rossmaessler,

1836

Unio fusculus Rossmaessler, 1836
Unio squamosus Charpentier, 1837
Unio gangraenosus J. F. Schmidt,

1840

Unio pruinosus J. F. Schmidt, 1840
Unio nessorhynchus Ktister, 1854
Unio heldii Kiister, 1854
Unio bosnensis von Moellendorff,

1873

Unio phaseolus Held in Kobelt, 1879
Unio neocomensis Drouet, 1881
Unio croaticus Drouet, 1881
Unio striatulus Drouet, 1882

Unio truncatulus Drouet, 1883
Unio rivalis Drouet, 1884
Unio serbicus Drouet, 1884
Unio savensis Drouet, 1884
Unio dokici Drouet, 1884
Unio ondovensis Hazay, 1885
Unio albensis Hazay, 1885
Unio subrobustus Servain, 1885
Unio duregicus Servain, 1885
Unio sandriopsis Servain, 1885
Unio ostiorum Servain, 1885
Unio turicus Servain, 1885
Unio tiguricus Servain, 1885
Unio travnicensis Kobelt, 1910
Unio brandisi Kobelt, 1910
Unio consentaneus hohenwarti Ko-
belt, 1910

Unio langi Kobelt, 1910
Unio (consentaneus var.?) gallen-

steini Kobelt, 1910
Unio consentaneus subterminalis

Kobelt, 1910

Unio batavus elongatus Clessin, 1910
Unio consentaneus thayacus Kobelt,

1910
Unio consentaneus biaudeti Kobelt,

1911

Unio consentaneus bodamicus Ko-
belt, 1911
Unio consentaneus dilatatus Kobelt,

1911

Unio consentaneus abnobae Kobelt,
1911

May hybridize with the Atlantic C7. c. batavus.
Rhone basin to and inclusive of Lake Geneva; Swiss Rhine and
tributaries; Danube system; Galicia; Wardar River.

Unio crassus batavus Maton and

Unio nana Lamarck, 1819
Unio riparia C. Pfeiffer, 1821
Unio arcuatus Bouch.-Chant., 1838
Unio droueti Dupuy, 1849
Unio courtillieri Hatteman, 1859
Unio batavus var. minimus Joannis,

1859
Unio batavus var. baraceus Joannis,

1859

Unio lamboltei Malzine, 1867
Unio subtilis Drouet, 1879
Unio ligericus Bourguignat, 1882
Unio dubisianus Coutagne, 1882
Unio sequanicus Coutagne, 1882
Unio rayi Bourguignat, 1882
Unio pilloti Bourguignat, 1882
Unio crassatellus Bourguignat in

Locard, 1882
Unio socardianus Bourguignat in

Locard, 1882
Unio locardianus Bourguignat in

Locard, 1882

Rackett, 1807

Unio lagnysiacus Locard, 1882
Unio macrorhynchus Bourguignat,

1882

Unio melas Coutagne, 1882
Unio dubisianopsis Locard, 1882
Unio carantoni Coutagne, 1882
Unio minutulus Ray, 1882
Unio berthellini Bourguignat, 1882
Unio andegavensis Servain, 1882
Unio cyprinorum Berthier, 1882
Unio matronicus Bourguignat, 1882
Unio septentrionalis Bourguignat,

1882

Unio oxyrhynchus Breviere, 1882
Unio marcellinus Berthier, 1882
Unio potamius Bourguignat, 1882
Unio feliciani Bourguignat, 1882
Unio arenarum Bourguignat, 1882
Unio danemorae Bourguignat, 1882
Unio brevieri Bourguignat, 1882
Unio tumidiformis Castro, 1885
Unio sadoicus Castro, 1885

1940

PALEARCTIC UNIONIDS — HAAS

131

Unio macropygus Castro, 1885
Unio eupygus Castro, 1885
Unio batavellus Letourneux, 1885
Unio batavus var. taunica Kobelt,

1886

Unio baeticus Kobelt, 1887
Unio carcasinus Souverbie, 1887
Unio besnardianus Servain, 1888
Unio baudoni Folin, 1888
Unio crassulus Drouet, 1888
Unio vegesackensis Servain, 1888
Unio visurgisinus Servain, 1888
Unio badiellus Drouet, 1888
Unio catalaunicus Coutagne, 1889
Unio zoasthenicus Locard, 1889
Unio jurianus Locard, 1889
Unio subamnicus Locard, 1889
Unio aturicus Locard, 1889
Unio lemotheuxi Servain, 1889
Unio orbus Locard, 1889
Unio andeliacus Bourguignat, 1889
Unio vallieriacus Bourguignat, 1889
Unio nubilus Locard, 1889
Unio diptychus Surrault, 1889
Unio ingrandiensis Surrault, 1889
Unio materniacus Locard, 1889
Unio melantatus Bourguignat, 1889
Unio balbignyanus Bourguignat,

1889

Unio occidentalis Bourguignat, 1889
Unio stygnus Locard, 1889
Unio bouchardi Bourguignat, 1889
Unio passavanti Bourguignat, 1889
Unio surraulti Servain, 1889
Unio adonus Servain, 1889
Unio camonti Bourguignat, 1889

Unio hattmanni Bourguignat, 1889
Unio seneauxi Bourguignat, 1889
Unio visurgicus Servain, 1889
Unio carioliensis Pac6me, 1889
Unio mariae Pac6me, 1889
Unio nicolloni Locard, 1889
Unio euthymeanus Locard, 1889
Unio ignariformis Bourguignat,

1889

Unio ignari Bourguignat, 1889
Unio manculus Locard, 1889
Unio perroudi Locard, 1889
Unio scotinus Locard, 1889
Unio redactus Locard, 1889
Unio conimbricus Kobelt, 1893
Unio gliscerus Locard, 1893
Unio ampullaceus Locard, 1893
Unio chardoni Bourguignat in

Locard, 1893

Unio lancelevei Locard, 1893
Unio hassiae Haas, 1908
Unio pseudocrassus Haas, 1909
Unio batavus sabulosus Haas, 1910
Unio batavus badensis Haas, 1910
Unio batavus palatinus Haas, 1911
Unio batavus hexameri Haas, 1911
Unio batavus probavaricus Haas,

1911
Unio batavus distinguendus C.

Boettger, 1912
Unio batavus navensis C. Boettger,

1912

Unio batavus kobeltianus Haas, 1913
f Unio cantianus Kennard and Wood-
ward, 1924

This list is a first and incomplete attempt to gather the synonyms
of this highly variable race.

Atlantic Iberian Peninsula; Atlantic France; western Germany,
not including the Weser; subfossil in England.

Unio crassus carneus Kiister, 1848

Unio luxurians Kiister, 1848

Montenegro; Albania.

Unio brevirostris Kiister, 1848

Unio crassus jonicus Drouet, 1879

Unio helenae Kobelt, 1893

Western Greece; isles of Corfu and Levkas.
Unio crassus gontieri Bourguignat, 1856

Unio colchicus Drouet, 1881 Unio stepanovi Drouet, 1881

Unio sieversi Drouet, 1881
Unio stevenianus Drouet, 1881
Unio mingrelicus Drouet, 1881
Unio raddei Drouet, 1881
Unio araxenus Drouet, 1881

Unio raddei var. kutaisanus Kobelt,

1886

Unio crassus okae Kobelt, 1911
Unio kungurensis Kobelt, 1912

132 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Unio kungurensis var. irenjensis

Kobelt, 1912
Unio kungurensis var. sylvensis

Kobelt, 1912
Unio kungurensis var. chlebnikowi

Kobelt, 1912

Unio armeniacus Kobelt, 1912
Unio roseni Kobelt, 1912
Unio gregorii Kobelt, 1912
Unio subbatavus Kobelt, 1913
Unio mingrelicus var. steveniani-
formis Shadin, 1938

This subspecies has been cited as stevenianus Krynicki, 1837,
but this name remained a nomen nudum until 1881, and gontieri,
dating from 1856, has priority.1

Caucasus; Basin of River Kama; White Russia.
Unio crassus bruguierianus Bourguignat, 1853

Unio orientalis Bourguignat, 1852

(not Lea, 1840)

Unio prusii Bourguignat, 1853
Unio vescoi Bourguignat, 1856
Unio schwerzenbachi Bourguignat,

1856

Unio turcicus Kuster, 1862
Unio kotschyi Kuster, 1862
Unio damascensis Lea, 1863
Unio orontesensis Lea, 1863
Unio syriacus Lea, 1863
Unio orphaensis Lea, 1864
Unio mardinensis Lea, 1864
Unio kullethensis Lea, 1864
Unio vicarius Westerlund, 1879
Unio desectus Westerlund, 1879
Unio heldreichi O. Boettger, 1886

Eastern Greece; Asia Minor.

Unio thiesseae Drouet, 1892
Unio scissus Drouet, 1893
Unio pseudonymus Simpson, 1900
(?)£7rao mpdiola Preston, 1912
Unio specialis Kobelt, 1913
Unio ancyrensis Kobelt, 1913
Unio ancyrensis var. louisei Kobelt,

1913
Unio orphaensis var. ehrmanni

Kobelt, 1913

Unio diarbekrianus Kobelt, 1913
Unio bitlisensis Kobelt, 1913
Unio ciconius Kobelt, 1913
Unio medicus Kobelt, 1913
Unio thospiensis Kobelt, 1915
Unio (Rhombunio) wenzi Kobelt,

1915

Unio crassus mongolicus Middendorff, 1851

Known only from Middendorff's original description. Preston
(1912) cited it from the upper Indus River, but this information does
not seem to be reliable.

Amur River System, northeastern Asia.

The three genera which now follow, i.e. Leguminaia, Pseudo-
dontopsis and Microcondylaea, are only intuitively grouped with the
Unionines, since anatomical evidence is not yet available. The
three genera cited above are close relatives of the genus Pseudodon
Gould, which is now restricted to southeastern Asia, though it ranged
much farther westward in late Tertiary time.

1 Unio steveniana Krynicki, Bull. soc. imp. nat. Moscou, 10, p. 59, 1837 (nomen
nudum); Siemaschko, Bull. soc. imp. nat. Moscou, 22, p. 126, 1847 (nomen nudum);
Unio gontieri Bourguignat, Amenites Malacologiques, 2, p. 33, pi. 4, figs. 1-4, 1856;
Unio stevenianus Drouet, Union. Russie Europ., p. 14, 1881 (first description of
species).

1940

PALEARCTIC UNIONIDS — HAAS

133

Genus Leguminaia Conrad, 1865

Leguminaia wheatleyi Lea, 1862

Monocondylaea mardinensis Lea,

1864

Leguminaia chantrei Locard, 1883
Leguminaia bourguignati Locard,

1883

Leguminaia naegelei Kobelt, 1913
Leguminaia graeteri Kobelt, 1913
Leguminaia nisibina Kobelt, 1913
Leguminaia haasi Kobelt, 1915

Iraq; Kara-su in Syria; Lake Antioch, Syria.

Leguminaia saulcyi Bourguignat, 1852

Unio michonii Bourguignat, 1852
Unio tripolitanus Bourguignat, 1852

Pseudodon chantrei Locard, 1883
Leguminaia locardi Simpson, 1900

Germain (1911) has created a new subgenus Pseudoleguminaia
for this species, which is characterized by oval shape, but I am
unwilling to recognize it.

Lake Antioch; rivers of Syrian littoral to the Nahr-el-Audja
near Jaffa in the south.

Genus Pseudodontopsis Kobelt, 1913
Pseudodontopsis euphraticus Bourguignat, 1852

Unio opperti Bourguignat, 1856
Pseudodon churchillianus Bourgui-
gnat, 1857

Monocondylaea rhomboidea Lea,
1859

Pseudodon pachyolenus Bourgui-
gnat, 1881 (nomen nudum)

Pseudodontopsis piestius Kobelt,
1913

Pseudodontopsis babylonicus Kobelt,
1913

Iraq; churchillianus is said to come from Konieh.

Genus Microcondylaea von Vest, 1866
Microcondylaea compressa Menke, 1830

(l)Anodonta uniopsis Lamarck,

1819; unidentifiable
Unio depressa C. Pfeiffer, 1825 (not

Lamarck, 1819)
Unio bonellii Ferussac in Ross-

maessler, 1835
Unio bonellii var. curvata Ross-

maessler, 1835
Unio moreleti Droue't, 1879
Microcondylus crassus Droue't, 1879
Microcondylus gibbosus Droue't,

1879
Microcondylus squamosus Droue't,

1879
Microcondylus truncatus Droue't,

1879

Microcondylaea servaini Bourgui-
gnat, 1883

Microcondylaea doriae Bourgui-
gnat, 1883

Microcondylaea gestroi Bourguignat,
1883

Leguminaia pedemontana Pollonera,
1889

Leguminaia craverii Pollonera,
1889

Microcondylaea bonellii f. parisii
Kobelt, 1913

Microcondylaea bonellii f. levicensis
Kobelt, 1913

Pseudanodonta depressa var. augusti
Graziadei, 1933

Po basin; Adige basin; littoral rivers east of Adige to and com-
prising the Isonzo.

134 FIELD MUSEUM OF NATURAL HISTORY— ZOOLOGY, VOL. 24

Subfamily Quadrulinae

This holarctic subfamily has only one representative in the
palearctic region at the present time, Psilunio Stefanescu; but in
the Pliocene there were more quadruline genera in Europe, which
were closely related to the modern east-Asiatic genera Cuneopsis
Simpson and Lamprotula Simpson. The center of distribution of
the quadrulines is North America.

The quadruline nature of Psilunio was proved by the study of
its anatomy (Haas, 1920, 1924).

Genus Psilunio Stefanescu, 1896

Rhytia Stefanescu, 1896 Rhombunio Germain, 1911

Psilunio acarnanicus Kobelt, 1879

Unio acarnanicus var. messenicus
Westerlund, 1879

Unio pamisinus Drouet, 1894

Western Greece, in Lake Vrachori; in Messenia and in the
Eurotas River.

Psilunio littoralis Lamarck, 1801
Psilunio littoralis littoralis Lamarck, 1801

Unio brevialis Lamarck, 1819
Unio subteiragoniis Michaud, 1831
Unio draparnaldii Deshayes, 1831
Unio pianensis Farines, 1833
Unio cuneatus Jacquemin, 1835
Unio rotundatus Mauduyt, 1839
Unio barraudi Bonhomme, 1840
Unio bigerrensis Millet, 1843
Unio moulinsianus Dupuy, 1850
Unio astierianus Dupuy, 1850
Unio rhomboideus Moquin-Tandon,

1855

Unio unduliferus Kiister, 1861
Unio subreniformis Bourguignat,

1863
ft/mo hippopotami Bourguignat,

1869

Unio rathymus Bourguignat in

Locard, 1882

Unio gandiensis Drouet, 1888
Unio pacomei Bourguignat in

Locard, 1889
Unio circulus Bourguignat in

Locard, 1893
Unio sphaericus Bourguignat in

Locard, 1893

Unio ovuliformis Locard, 1893
Unio rhysopygus Drouet, 1894
Unio circinatus Drouet, 1894
Unio mauduyti Germain, 1897
Unio littoralis var. taginus Kobelt,

1903

ft/mo kinkelini Haas, 1910
Unio batavus calalonicus Haas, 1921

Subfossil in the Rhine basin and in South England; recent in
France, Iberian Peninsula with the exception of the southern part.

Psilunio littoralis umbonatus Rossmaessler, 1844

(l)Unio incurvus Lea, 1831; said to
have come from Gibraltar, un-
identifiable

Unio hispalensis Kobelt, 1887

Southern part of Iberian Peninsula.

Unio colder oni Kobelt, 1887
Unio calderoni var. salvadori Wes-
terlund, 1892

1940

PALEARCTIC UNIONIDS— HAAS

135

Psilunio littoralis komarowi 0. Boettger, 1880
Armenia.

Psilunio littoralis semirugatus Lamarck, 1819

Unio rothi Bourguignat, 1863
Unio emesaensis Lea, 1864
Unio simonis Tristram, 1865
Unio timius Bourguignat in Locard,

1883

Unio galilaei Locard, 1883
Unio luynesi Locard, 1883
Unio rhomboidopsis Locard, 1883
Unio tinctus Drouet, 1893
Unio rollei Kobelt, 1895

Syria; Jordan basin in Palestine.

Unio graeteri Kobelt, 1913
Unio babensis Kobelt, 1913
Unio naegelei Kobelt, 1913
Unio beroeus Kobelt, 1913
Unio halepensis Kobelt, 1913
Unio halepensis var. cazioti Kobelt,

1913

Unio corbiculiformis Kobelt, 1913
Unio chinnerethensis Preston, 1913
Unio salamboana Pallary, 1929

Psilunio littoralis homsensis Lea, 1864

Unio episcopalis Tristram, 1865 Margaritana syriaca Pallary, 1929

Unio barroisi Drouet, 1893

"Margaritana" syriaca is an unmistakable Psilunio, in spite of
its elongated, ear-shaped shell; this becomes evident by the umbonal
sculpture, the hinge, the roundish juvenile shell, the deep umbonal
cavity, and other characters, and Pallary (1929) quite inadequately
arranged it with Margaritana (=Margaritifera). It is by no means
impossible that homsensis Lea can not be retained as a separate sub-
species, but has to be united with semirugatus, of which it may be a
rather extraordinarily elongated form. Unio tinctus Drouet, liv-
ing together with homsensis in the Orontes River, combines a rounded,
typical Psilunio shape of the shell with the purple nacre, by which
homsensis is characterized. Unio modiola Preston, 1912, which I
place with Unio crassus bruguierianus Bourguignat, may as well
belong to homsensis; its description is too poor to make possible a
decision on this point.

Orontes River, Syria.

Psilunio littoralis delesserti Bourguignat, 1853

Unio ferussacianus Lea, 1868
Unio wagneri Kobelt, 1895
Unio tracheae Kobelt, 1895
Unio lycicus Kobelt, 1895

Littoral rivers of Palestine.

Unio deschampsi Kobelt, 1913
Unio langloisi Kobelt, 1913
Unio abrus Kobelt, 1913
Unio blanchianus Kobelt, 1913

Psilunio littoralis fellmanni Deshayes, 1847

Unio ravoisieri Deshayes, 1847
Unio moreleti Deshayes, 1847
Unio unduliferus Kiister, 1861
Unio maccarthyanus Bourguignat,
1866

Unio mauritanicus Bourguignat,

1868

Unio ksibianus Mousson, 1873
Unio jolyanus Bourguignat in

Kobelt, 1886

136 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Unio zenaticus Letourneux in Bour- Unio rhomboideus var. bucheti Pal-

guignat, 1887 lary, 1921

Unio rouirei Bourguignat, 1887 Unio tifleticus Pallary, 1927

Unio marteli Pallary, 1918 Rhombunio asananus Pallary, 1928

Unio unduliferus Kiister (Martini-Chemnitz, 9, pt. 2, Unio,
p. 162, pi. 46, fig. 4, 1861) described as probably of South American
origin, certainly belongs here.

North Africa.

Subfamily Anodontinae

Contains the holarctic genus Anodonta; also Pseudanodonta,
which is a close relative of the Asiatic genus Pilsbryoconcha, and
Gabillotia, a European endemism.

Genus Anodonta Lamarck, 1799
Anodonta cygnea Group

Anodonta cygnea Linnaeus, 1758

Under this name I combine all the palearctic anodontas, with
one exception, which will be mentioned below. The enormous
plasticity and variability of the Anodonta shell have led to the pro-
posal of many hundreds of specific names, in which not only Bour-
guignat and his school have shared; for even conscientious scientists
of the other camp have again and again been induced by apparently
characteristically shaped pond-mussels, to propose new names,
being convinced of having separable species or, at least, local races
before them.

There is only one — or, according to the most modern researches,
two — fundamental species of Anodonta in the palearctic region, to
which all the many different-looking "species" that have been
described can be reduced. I myself, when beginning my studies on
unionids in 1910, was inclined to recognize two such fundamental
species, the broader Anodonta cygnea and the more elongated
Anodonta cellensis Schroeter. This point of view I have meantime
abandoned; it is at present maintained by Franz in Jena, and by
his school, who recognize Anodonta cygnea and Anodonta piscinalis
Nilsson as fundamental species. I am of the opinion that all the
palearctic anodontas belong to one unique species, with the excep-
tion above indicated. It is worth mentioning that Clessin also
arrived at the same conviction; he described an Anodonta mutabilis
which was to comprise all the palearctic pond-mussels, including
even their typical Linnaean species, cygnea.

1940

PALEARCTIC UNIONIDS— HAAS

137

Anodonta cygnea, as it is here understood, lives throughout the
palearctic region. In northern and central Europe, it is a relatively
common species, but becomes less so in the Mediterranean basin,
is scarce in North Africa and in Asia Minor, and is only locally
known from central Asia; for instance, from Samarkand (samarkan-
densis Kobelt), Buchara (sogdiana Kobelt, bactriana Rolle), the
Seisan-Nor (seisanensis Kobelt), the Lena River (lenae Shadin), Lake
Baikal (sorensiana Dybowsky, sorica Dybowsky) and its tributary,
the Selenga River (nova Dybowsky, selengensis Dybowsky).

Pallary has described (1933) a subgenus Euphrata, based upon
Anodonta bahlikiana (nomen nudum) from Mesopotamia, and Be'de'
has created (1932) another subgenus, Liouvillea, which rests on his
Moroccan species pallaryi and theryi. Not having seen them, I
cannot form a judgment of them; but knowing Pallary 's tendency
to publish new names for merely phaenotypic mutations, I rather
strongly suspect that both subgenera are founded upon environ-
mentally shaped and only apparently separable forms of Ano-
donta cygnea.

As I am unwilling to give up this point of view, it is unnecessary
for me to give a complete synonymic list of Anodonta cygnea. The
anodontas contained in Westerlund's Fauna constitute such a list
up to 1890, and the following enumeration includes only such
"species" as have been described since that date.

Anodonta ataxia Baichere, 1891
Anodonta culoxiana Nicolas, 1891
Anodonta suevica Kobelt, 1891
Anodonta borealis Kobelt, 1891
Anodonta recurmrostris Kobelt, 1892
Anodonta gallensteini Kobelt, 1892
Anodonta graeca Drouet, 1892
Anodonta quelleneci Drouet, 1892
Anodonta lepida Drouet, 1892
Anodonta Ihotellerii Drouet, 1892
Anodonta discoidea Drouet, 1893
Anodonta baudoniana Drouet, 1893
Anodonta planulata Drouet, 1893
Anodonta viridiflava Drouet, 1893
Anodonta macella Drouet, 1893
Anodonta lanceolata Drouet, 1893
Anodonta limbata Drouet, 1893
Anodonta laevigata Drouet, 1893
Anodonta retziana Drouet, 1893
Anodonta gregalis Drouet, 1893
Anodonta aeneolina Drouet, 1893
Anodonta pulchella Drouet, 1893
Anodonta fuliginea Drouet, 1893
Anodonta ervica Kobelt, 1894
Anodonta latirostris Drouet, 1894
Anodonta mollis Drouet, 1894
Anodonta adusta Drouet, 1894

Anodonta glaucina Drouet, 1894
Anodonta prasina Drouet, 1894
Anodonta nobilis Drouet, 1894
Anodonta bicolor Drouet, 1894
Anodonta valentina Drouet, 1894
Anodonta emacerata Drouet, 1894
Anodonta cilicica Kobelt, 1895
Anodonta simulans Drouet, 1895
Anodonta sogdiana Kobelt, 1896
Anodonta bactriana Rolle, 1897
Anodonta fedderseni Westerlund,

1898

Anodonta goesi Westerlund, 1898
Anodonta anura Westerlund, 1898
Anodonta apala Locard, 1899
Anodonta pelophila Locard, 1899
Anodonta simoesi Locard, 1899
Anodonta portensis Locard, 1899
Anodonta leprosa Locard, 1899
Anodonta retteri Drouet, 1899
Anodonta chivoti Germain, 1904
Anodonta bouvieri Rochebrune, 1904
Anodonta falcata var. fluviatica

Rosen, 1905
Anodonta (piscinalis var.) portulana

Kobelt, 1908

138 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24

Anodonta piscinalis transsylvanica Anodonta jesseni Schlesch, 1926

Haas, 1911 Anodonta (Liouvillea) pallaryi Bede,

Anodonta seisanensis Kobelt, 1912 1932

Anodonta anatina var. circularis Anodonta (Liouvillea) theryi Bede,

Hilbert, 1912 1932

Anodonta (cyrea var.) samarkanden- Anodonta (Euphrata) bahlikiana

sis Kobelt, 1913 Pallary, 1933 (nomen nudum)

Anodonta piscinalis var. sorica Anodonta cygnea var. armenica

Dybowsky, 1913 Ovtshinnikov, 1935

Anodonta complanata var. sorensi- Anodonta anatina var. petshorica

ana Dybowsky, 1913 Shadin, 1938

Anodonta nova Dybowsky, 1913 Anodonta anatina var. lenae Shadin,

Anodonta cellensis var. selengensis 1938

Dybowsky, 1913 Anodonta piscinalis var. volgensis

Anodonta waterstoni Tomlin, 1923 Shadin, 1938

The genus Cristaria Schumacher, so widely distributed in eastern
Asia, does not form part of the actual palearctic fauna. Neverthe-
less, some lacustrine European Anodonta cygnea which have acquired
the characteristic shell features of Cristaria, i.e., a symphynote shell
and a kind of hinge composed of claustra, may be considered as con-
vergent forms; so the Danubian Anodonta cygnea, upon which Bour-
guignat (1881) based his genus Colletopterum, those from Lake
Balaton (Hungary), Lake Doiran (Macedonia), and from the Albu-
fera de Valencia (Spain). But it must be emphasized that the shell
features we just mentioned are genotypically anchored in Cristaria
and only phaenotypically developed in lacustrine European Ano-
donta cygnea.

Anodonta woodiana Group

With this East-Asiatic species, to which the group of the North
American Anodonta grandis Say, 1829, belongs, I place the following
palearctic species:

Anodonta vescoiana Bourguignat, 1857

Anodonta vescoiana var. mesopo- Anodonta schlaeflii Mousson, 1874

tamica Mousson, 1874

Iraq. If Bourguignat's information, that his Anodonta vescoiana
originated from Konieh, Asia Minor, is correct, then his Anodonta
apollonica and Anodonta taurica, both coming from the Lake of
Apollonia, Asia Minor, may belong to the woodiana group.

Genus Pseudanodonta Bourguignat, 1877

Upon this genus opinions still differ widely. While some scien-
tists do not regard it even as a subgenus of Anodonta, I consider it
as a separable genus, well characterized by its peculiar shape and
by the features of its glochidium; it is most closely related to the
genus Pilsbryoconcha Simpson of southeastern Asia. Quite in con-

1940

PALEARCTIC UNIONIDS— HAAS

139

trast with Anodonta, Pseudanodonta forms local races, but it is so
variable that there is as yet no agreement about these local races
or even about its fundamental species. Thus the arrangement given
below will soon need amendment.

Pseudanodonta elongata Holandre, 1836

Anodonta grateloupiana Gassies,
1849

Anodonta grateloupiana var. globosa
Gassies, 1849

Anodonta normandi Dupuy, 1849

Anodonta jobae Dupuy, 1849

Anodonta siliqua Ktister, 1852

Pseudanodonta rayi Bourguignat,
1881

Pseudanodonta ligurica Bourgui-
gnat, 1881

Anodonta dorsuosa Drouet, 1881

Pseudanodonta locardi Locard, 1882

Pseudanodonta ararisana Locard,
1882

Pseudanodonta servaini Bocard,
1885

Anodonta pachyproktus Borcher-
ding, 1889

Anodonta fusiformis Borcherding,
1889

Anodonta microptera Borcherding,
1889

Pseudanodonta rothomagensis
Locard, 1890

Pseudanodonta nantelica Bourgui-
gnat, 1890

Pseudanodonta pechaudi Bourgui-
gnat, 1890

Pseudanodonta imperialis Servain,
1890

Pseudanodonta arnouldi Pacdme,
1890

Pseudanodonta isarana Bourgui-
gnat, 1890

Pseudanodonta mongazonae Bour-
guignat, 1890

Pseudanodonta lacustris Servain,
1890

Pseudanodonta rivalis Bourguignat,
1890

Pseudanodonta septentrionalis
Locard, 1890

Pseudanodonta aploa Bourguignat,
1890

Pseudanodonta euthymei Pacdme,
1890

Pseudanodonta morini Locard, 1890

Pseudanodonta cazioti Bourguignat,
1890

Pseudanodonta pacomei Bourgui-
gnat, 1890

Pseudanodonta trivurtina Bourgui-
gnat, 1890

Pseudanodonta brebissoni Locard,
1890

Anodonta albica Drouet, 1893

Anodonta (Pseudanodonta) acutalis
Drouet in Westerlund, 1894

Pseudanodonta dumasi Locard, 1894

Pseudanodonta beryacensis Dumas,
1894

Pseudanodonta limosina Dumas,
1895

Pseudanodonta ocddentalis Cou-
tagne, 1895

Pseudanodonta nicarica Haas, 1908

Atlantic western Europe to and comprising the Weser in the
East; England; France approximately to the Garonne in the south.

Pseudanodonta complanata Rossmaessler, 1835

Anodonta klettii Rossmaessler, 1835
Pseudanodonta nordenskioldii Bour-
guignat, 1881

Pseudanodonta complanata silesiaca

Kobelt, 1911
Pseudanodonta maelarensis Kobelt,

1911

Pseudanodonta borealis Kobelt, 1890, from Newa River, Russia,
which occasionally is cited as belonging here, is based upon a mis-
understood long and slender form of Anodonta cygnea Linnaeus.

Sweden; Atlantic Central Europe from the Elbe in the west to
Finland in the east.

140 FIELD MUSEUM OF NATURAL HISTORY — ZOOLOGY, VOL. 24
Pseudanodonta middendorffi Siemaschko, 1848
Pseudanodonta middendorffi middendorffi Siemaschko, 1848

Anodonta complanata jijiana Nico- Anodonta complanata sobriewskii

dim, 1909 Rosen, 1925

Pseudanodonta euxina Rossmaess-

ler in Kobelt, 1911

Tributaries of the Black Sea, east of the Danube.
Pseudanodonta middendorffi compacta Zelebor, 1851

Alasmodonta penchinati Bourgui- Pseudanodonta pantiti Bourgui-

gnat, 1870 gnat, 1881

Alasmodonta berlandi Bourguignat, Pseudanodonta scrupea Bourgui-

1870 gnat, 1881

Pseudanodonta praedara Bourgui- Pseudanodonta rossmaessleri Bour-

gnat, 1881 guignat, 1881

Pseudanodonta letourneuxi Bour- Pseudanodonta ellipsiformis Bour-
guignat, 1881 guignat, 1881

Pseudanodonta danubialis Bourgui- Pseudanodonta savensis Kobelt,

gnat, 1881 1911

Pseudanodonta mecyna Bourgui- Pseudanodonta compacta kiisteri

gnat, 1881 Haas, 1913

Pseudanodonta compacta kiisteri apparently hybridizes with the
Atlantic P. elongata where their ranges overlap in the Regnitz basin,
tributary to the Rhine via the Main (Haas and Schwarz, 1913).

Basin of the Danube.

Genus Gabillotia Servain, 1890

This genus is endemic in the palearctic region and is most closely
related to Anodonta.

Gabillotia pseudodopsis Locard, 1883

Gabillotia locardi Servain, 1890

Known only from Lake Antioch in Syria.

APPENDIX

Unionids which have erroneously been attributed to the palearc-
tic region:

Unio bagdadensis Bourguignat, 1852. — Said to come from

Baghdad, but is a synonym of Caelatura egyptiaca Caillaud

from the Nile.
Unio eucyphus Bourguignat, 1857. — Believed to have been

found in the Skamander River, northern Asia Minor, but

equally identical with Caelatura egyptiaca Caillaud from the

Nile.

1940 PALEARCTIC UNIONIDS — HAAS 141

Unio bythinicus Kobelt, 1893. — Not from Asia Minor, as believed
by Kobelt, being nothing but Elongaria orientalis Lea from
Java.

Unio lapidosus Kobelt, 1893. — Supposed to be from the
Euphrates River; it isLampsilis (Disconaias) discus Lea from
North Mexico.

Unio wolwichi Morelet, 1845. — This species, which was said to
have been found at the banks of the River Tajo in Portugal,
has been proved to be a synonym of Diplodon parallelipipedon
Lea from the Rio de la Plata.

REFERENCES

BEDS, P.

1932. Le genre "Anodonta" au Maroc. Bull. Soc. Sci. Nat. Maroc, 12, pp.
223-227.

BOURGUIGNAT, J. R.

1881. Materiaux pour servir a 1'histoire des Mollusques Acephales du systeme
europeen. 387 pp. Poissy.

GERMAIN, L.

1911. Mollusques terrestres et fluviatiles de 1'Asie anterieure. Bull. Mus.
Hist. Nat. Paris, 17, pp. 63-67.

HAAS, F.

1910. Die Najadenfauna des Oberrheins vom Diluvium bis zur Jetztzeit.

Abh. Senck. Nat. Ges., 32, pp. 143-176, 3 pis.
1920. Die Gattung Rhombunio, ihre Anatomie und Stellung im System.

Senckenbergiana, 2, pp. 70-80.
1924. Anatomische Untersuchungen an europaischen Najaden. I. Arch. Moll.

Kunde, 55, pp. 66-82, pis. 4-5.

— and SCHWARZ, E.

1913. Die Unioniden des Gebietes zwischen Main und deutscher Donau in
tiergeographischer und biologischer Hinsicht. Abh. Akad. Wiss. Mlinchen,
math.-phys. Kl., 26, Abh. 7, 34 pp., 4 pis.

PALLARY, P.

1929. Premiere addition a la faune malacologique de la Syrie. Mem. Inst.
Egypt., 12, pp. 1-43, pis. 1-3.

1933. Resultats generaux d'une prospection malacologique effectuee en Syrie
de 1929 a 1932. Bull. Mus. Hist. Nat. Paris, (2), 5, pp. 148-154.

PR ASH AD, B.

1919. On the Generic Position of Some Asiatic Unionidae. Rec. Ind. Mus.
Calcutta, 16, pp. 403-411.

PRESTON, H. B.

1912. Catalogue of the Asiatic Naiades in the Collection of the Indian Mu-
seum, Calcutta, with Descriptions of New Species. Rec. Ind. Mus. Calcutta,
7, pp. 279-308, pi. 8.

WESTERLUND, C. A.

1890. Fauna der in der palaarctischen Region lebenden Binnenconchylien.
VII: Malacozoa Acephala. 319 pp. Berlin.

UNIVERSITY OF ILLINOIS-URBANA